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34200.9684Heat-shock-increased survival to far-UV radiation in Escherichia coli is wavelength dependent. Heat-shock-induced resistance to far-UV (FUV) radiation was studied in Escherichia coli. The induction of FUV resistance was shown to be dependent on the products of the genes uvrA and polA in bacteria irradiated at 254 nm. Heat shock increased the resistance to 280 nm radiation in a uvrA6 recA13 mutant. Heat shock lowered the mutation frequency (reversion to tryptophan proficiency) in wild-type or uvrA strains irradiated at 254 nm. When these strains were irradiated at 280 nm, heat shock did not interfere with the mutation frequency in the wild-type strain, but greatly enhanced mutations in the uvrA mutant. After heat-shock treatment, the wild-type strain irradiated at 254 nm showed increased DNA degradation, indicating enhanced repair activity. However, heat shock did not stimulate SOS repair triggered by FUV. An increased survival of bacteriophages irradiated with FUV and inoculated into heat-shock-treated bacteria was not detected. The possibility that heat shock enhances excision repair activity in a wavelength-dependent manner is discussed.19948176549
811210.9666Fate of antibiotic resistance bacteria and genes during enhanced anaerobic digestion of sewage sludge by microwave pretreatment. The fate of antibiotic resistant bacteria (ARB) and antibiotic resistance genes (ARGs) were investigated during the sludge anaerobic digestion (AD) with microwave-acid (MW-H), microwave (MW) and microwave-H2O2-alkaline (MW-H2O2) pretreatments. Results showed that combined MW pretreatment especially for the MW-H pretreatment could efficiently reduce the ARB concentration, and most ARG concentrations tended to attenuate during the pretreatment. The subsequent AD showed evident removal of the ARB, but most ARGs were enriched after AD. Only the concentration of tetX kept continuous declination during the whole sludge treatment. The total ARGs concentration showed significant correlation with 16S rRNA during the pretreatment and AD. Compared with unpretreated sludge, the AD of MW and MW-H2O2 pretreated sludge presented slightly better ARB and ARGs reduction efficiency.201626970692
34020.9663Study of MFD-type repair in locus determining resistance of Escherichia coli to streptomycin. The yield of induced mutations to streptomycin resistance (Str) in E. coli, UV-irradiated and temporarily incubated in liquid medium not permitting protein synthesis, depends upon the conditions of preirradiation growth and preirradiation treatment of the bacteria, i.e. on their physiological state at the moment of irradiation. This fact is not readily reconciled with a model postulating mutation production in the structural genes of E. coli during excision repair. A preferred explanation is offered, based on the assumption that the efficiency of mutagenesis at the rpsL (strA) locus is determined by interference of antimutagenic (generalized excision repair and MFD) and promutagenic (mutation fixation of excision repair) events. The participation of macromolecular syntheses in Str mutation fixation is suggested.19863537780
811730.9660Composting of oxytetracycline fermentation residue in combination with hydrothermal pretreatment for reducing antibiotic resistance genes enrichment. Hydrothermal pretreatment can efficiently remove the residual antibiotics in oxytetracycline fermentation residue (OFR), but its effect on antibiotic resistance genes (ARGs) during composting remains unclear. This study compared the shifts in bacterial community and evolutions in ARGs and integrons during different composting processes of OFRs with and without hydrothermal pretreatment. The results demonstrated that hydrothermal pretreatment increased the bacterial alpha diversity at the initial phase, and increased the relative abundances of Proteobacteria and Actinobacteria but decreased that of Bacteroidetes at the final phase by inactivating mycelia and removing residual oxytetracycline. Composting process inevitably elevated the abundance and relative abundance of ARGs. However, the increase in ARGs was significantly reduced by hydrothermal pretreatment, because the removal of oxytetracycline decreased their potential host bacteria and inhibited their horizontal gene transfer. The results demonstrated that hydrothermal pretreatment is an efficient strategy to reduce the enrichment of ARGs during the OFR composting.202033099099
848540.9659Nonsteroidal anti-inflammatory drug diclofenac accelerates the emergence of antibiotic resistance via mutagenesis. Overuse of antimicrobial agents are generally considered to be a key factor in the occurrence of antibiotic resistance bacteria (ARB). Nevertheless, it is unclear whether ARB can be induced by non-antibiotic chemicals such as nonsteroidal anti-inflammatory drug (NSAID). Thus, the objective of this study is to investigate whether NSAID diclofenac (DCF) promote the emergence of antibiotic resistance in Escherichia coli K12 MG1655. Our results suggested that DCF induced the occurrence of ARB which showed hereditary stability of resistance. Meanwhile, gene variation was identified on chromosome of the ARB, and DCF can cause bacterial oxidative stress and SOS response. Subsequently, transcriptional levels of antioxidant (soxS, sodA, sodC, gor, katG, ahpF) and SOS (recA, lexA, uvrA, uvrB, ruvA, ruvB, dinB, umuC, polB) system-related genes were enhanced. However, the expression of related genes cannot be increased in high-dosage treatment compared with low-dosage samples because of cytotoxicity and cellular damage. Simultaneously, high-dosage DCF decreased the mutation frequency but enhanced the resistance of mutants. Our findings expand our knowledge of the promoting effect on the emergence of ARB caused by DCF. More attention and regulations should be given to these potential ecological and health risks for widespread DCF.202336958653
793550.9654Removal of antibiotic resistance genes by Cl(2)-UV process: Direct UV damage outweighs free radicals in effectiveness. Antibiotic resistance genes (ARGs) pose significant environmental health problems and have become a major global concern. This study investigated the efficacy and mechanism of the Cl(2)-UV process (chlorine followed by UV irradiation) for removing ARGs in various forms. The Cl(2)-UV process caused irreversible damage to nearly all ARB at typical disinfectant dosages. In solutions containing only extracellular ARGs (eARGs), the Cl₂-UV process achieved over 99.0 % degradation of eARGs. When both eARGs and intracellular ARGs (iARGs) were present, the process reached a 97.2 % removal rate for iARGs. While the abundance of eARGs initially increased due to the release of iARGs from lysed cells during pre-chlorination, subsequent UV irradiation rapidly degraded the released eARGs, restoring their abundance to near-initial levels by the end of the Cl₂-UV process. Analysis of the roles in degrading eARGs and iARGs during the Cl(2)-UV process revealed that UV, rather than free radicals, was the dominant factor causing ARG damage. Pre-chlorination enhanced direct UV damage to eARGs and iARGs by altering plasmid conformation and promoting efficient damage to high UV-absorbing cellular components. Furthermore, no further natural transformation of residual ARGs occurred following the Cl(2)-UV treatment. This study demonstrated strong evidence for the effectiveness of the Cl(2)-UV process in controlling antibiotic resistance.202540048777
33960.9651Multiple mechanisms of resistance to cisplatin toxicity in an Escherichia coli K12 mutant. The mechanisms underlying cellular resistance to the antitumor drug cis-diamminedichloro-platinum(II) (CDDP) were studied in Escherichia coli K12. A bacterial strain (MC4100/DDP) was selected from the MC4100 wild-type strain after growth for four cycles in CDDP. MC4100/DDP bacteria showed a high level of resistance and exhibited various modifications including (1) a decrease in drug uptake and platinum/DNA binding which only partly contributed to resistance, (2) an increase in glutathione content not involved in the resistant phenotype, (3) an increase in DNA repair capacity. Resistance was unmodified by introducing a uvrA mutation which neutralizes the excision-repair pathway. In contrast, it was abolished by deletion of the recA gene which abolishes recombination and SOS repair but also by a mutation in the recA gene leading to RecA co-protease minus (no SOS induction). RecA protein was unchanged in MC4100/DDP but the expression of RecA-dependent gene(s) was required for CDDP resistance. The regulation of genes belonging to the SOS regulon was analysed in MC4100/DDP by monitoring the expression of sfiA and recA::lacZ gene fusions after UV irradiation. These gene fusions were derepressed faster and the optimal expression was obtained for a lower number of UV lesions in MC4100/DDP, suggesting a role of RecA co-protease activity in the mechanism of resistance to CDDP in this E. coli strain.19947974517
787270.9649Quaternary ammonium compounds promoted anoxic sludge granulation and altered propagation risk of intracellular and extracellular antibiotic resistance genes. Surfactants could influence sludge morphology and disinfectants were linked to antibiotic resistance genes (ARGs). Thus, the response of activated sludge and ARGs to long-term quaternary ammonium compounds (QACs) exposure required further investigation, which is a popular surfactant and disinfectant. Here, three sequencing batch reactors were fed with 5 mg/L most frequently detected QACs (dodecyl trimethyl ammonium chloride (ATMAC C12), dodecyl benzyl dimethyl ammonium chloride (BAC C12) and didodecyl dimethyl ammonium chloride (DADMAC C12)) for 180 d. The long-term inhibitory effect on denitrification ranked: DADMAC C12 > BAC C12 > ATMAC C12. Besides, obvious granular sludge promoted by the increase of α-Helix/(β-Sheet + Random coil) appeared in DADMAC C12 system. Moreover, intracellular ARGs increased when denitrification systems encountered QACs acutely but decreased in systems chronically exposed to QACs. Although replication and repair metabolism in ATMAC C12 system was higher, ATMAC C12 significantly promoted proliferation of extracellular ARGs. It was noteworthy that the propagation risk of extracellular ARGs in sludge increased significantly during sludge granulation process, and intracellular sul2 genes in sludge and water both increased with the granular diameter in DADMAC C12 system. The universal utilization of QACs may enhance antibiotic resistance of bacteria in wastewater treatment plants, deserving more attention.202336444811
785480.9648Removal of antibiotic resistant bacteria and plasmid-encoded antibiotic resistance genes in water by ozonation and electro-peroxone process. The electro-peroxone (EP) process is an electricity-based oxidation process enabled by electrochemically generating hydrogen peroxide (H(2)O(2)) from cathodic oxygen (O(2)) reduction during ozonation. In this study, the removal of antibiotic resistant bacteria (ARB) and plasmid-encoded antibiotic resistance genes (ARGs) during groundwater treatment by ozonation alone and the EP process was compared. Owing to the H(2)O(2)-promoted ozone (O(3)) conversion to hydroxyl radicals (•OH), higher •OH exposures, but lower O(3) exposures were obtained during the EP process than ozonation alone. This opposite change of O(3) and •OH exposures decreases the efficiency of ARB inactivation and ARG degradation moderately during the EP process compared with ozonation alone. These results suggest that regarding ARB inactivation and ARG degradation, the reduction of O(3) exposures may not be fully counterbalanced by the rise of •OH exposures when changing ozonation to the EP process. However, due to the rise of •OH exposure, plasmid DNA was more effectively cleaved to shorter fragments during the EP process than ozonation alone, which may decrease the risks of natural transformation of ARGs. These findings highlight that the influence of the EP process on ARB and ARG inactivation needs to be considered when implementing this process in water treatment.202336738938
792690.9647Microplastics Exacerbated Conjugative Transfer of Antibiotic Resistance Genes during Ultraviolet Disinfection: Highlighting Difference between Conventional and Biodegradable Ones. Microplastics (MPs) have been confirmed as a hotspot for antibiotic resistance genes (ARGs) in wastewater. However, the impact of MPs on the transfer of ARGs in wastewater treatment remains unclear. This study investigated the roles and mechanisms of conventional (polystyrene, PS) and biodegradable (polylactic acid, PLA) MPs in the conjugative transfer of ARGs during ultraviolet disinfection. The results showed that MPs significantly facilitated the conjugative transfer of ARGs compared with individual ultraviolet disinfection, and PSMPs exhibited higher facilitation than PLAMPs. The facilitation effects were attributed to light shielding and the production of reactive oxygen species (ROS) and nanoplastics from ultraviolet irradiation of MPs. The light shielding of MPs protected the bacteria and ARGs from ultraviolet inactivation. More importantly, ROS and nanoplastics generated from irradiated MPs induced intracellular oxidative stress on bacteria and further increased the cell membrane permeability and intercellular contact, ultimately enhancing the ARG exchange. The greater fragmentation of PSMPs than PLAMPs resulted in a higher intracellular oxidative stress and a stronger enhancement. This study highlights the concerns of conventional and biodegradable MPs associated with the transfer of ARGs during wastewater treatment, which provides new insights into the combined risks of MPs and ARGs in the environment.202539723446
7858100.9644Photocatalytic Reactive Ultrafiltration Membrane for Removal of Antibiotic Resistant Bacteria and Antibiotic Resistance Genes from Wastewater Effluent. Biological wastewater treatment is not effective in removal of antibiotic resistant bacteria (ARB) and antibiotic resistance genes (ARGs). In this study, we fabricated a photocatalytic reactive membrane by functionalizing polyvinylidene fluoride (PVDF) ultrafiltration (UF) membrane with titanium oxide (TiO(2)) nanoparticles for the removal of ARB and ARGs from a secondary wastewater effluent. The TiO(2)-modified PVDF membrane provided complete retention of ARB and effective photocatalytic degradation of ARGs and integrons. Specifically, the total removal efficiency of ARGs (i.e., plasmid-mediated floR, sul1, and sul2) with TiO(2)-modified PVDF membrane reached ∼98% after exposure to UV irradiation. Photocatalytic degradation of ARGs located in the genome was found to be more efficient than those located in plasmid. Excellent removal of integrons (i.e., intI1, intI2, and intI3) after UV treatment indicated that the horizontal transfer potential of ARGs was effectively controlled by the TiO(2) photocatalytic reaction. We also evaluated the antifouling properties of the TiO(2)-UF membrane to demonstrate its potential application in wastewater treatment.201829984583
7932110.9642How multi-walled carbon nanotubes in wastewater influence the fate of coexisting antibiotic resistant genes in the subsequent disinfection process. Wastewater treatment plants (WWTPs) are important hubs for the spread of antibiotic resistance genes (ARGs). Engineered nanoparticles, which was inevitably released to WWTPs, could change environmentally sensitive of antibiotic resistant bacteria (ARB). This would influence the fate of ARGs in subsequent disinfection process and consequent health risk. In this study, the ARGs fate of the effluent in conventional sodium hypochlorite (NaClO) disinfection process was investigated as multi-walled carbon nanotubes (MWCNTs) existed in sequencing batch reactor (SBR). The results showed the existence of MWCNTs in SBR could enhance the removal efficiency of intracellular 16S rRNA gene and intI1, extracellular intI1, sul2 and tetX in the effluent by NaClO. This is mainly due to the variation of bacterial physiological status, bacterial population structure and the activation of NaClO under the role of MWCNTs. MWCNTs in SBR could increase in membrane permeability of bacterial cells, which would be conducive to the penetration of chlorination to cytoplasm. MWCNTs in SBR also could change the bacterial population structure and induce the chlorine-sensitive bacteria; thus the potential hosts of ARGs in the effluent would be more easily inactivated by NaClO. Moreover, the residual MWCNTs in the effluent could activate NaClO to generate various free radical, which would enhance the oxidizing capacity of chlorination.202235500623
8491120.9642Hormesis-like effects of black phosphorus nanosheets on the spread of multiple antibiotic resistance genes. The production scalability and increasing demand for black phosphorus nanosheets (BPNSs) inevitably lead to environmental leakage. Although BPNSs' ecotoxicological effects have been demonstrated, their indirect health risks, such as inducing increased resistance in pathogenic bacteria, are often overlooked. This study explores the influence of BPNSs on the horizontal gene transfer of antibiotic resistance genes (ARGs) facilitated by the RP4 plasmid, which carries multiple resistance genes. The results indicated that BPNSs exhibited concentration-dependent hormesis-like effects on bacterial conjugation gene transfer. Specifically, at sub-inhibitory concentrations (0.0001-1 mg/L), BPNSs promoted both intra- and intergeneric conjugative transfer, demonstrating an initial increase followed by a decline, with transfer rates rising by 1.5-3.1-fold and 1.5-3.3-fold, respectively. BPNSs were found to induce reactive oxygen species (ROS) production, increase malondialdehyde levels, and trigger the SOS response, enhancing plasmid uptake. Additionally, BPNSs increased membrane permeability by forming pores and upregulating outer membrane porins (OMPs) genes. At higher BPNSs concentrations (0.1-1 mg/L), conjugative frequency was inhibited due to the disruption of the cellular antioxidant system and changes in the adsorption process. These findings underscore the influence of BPNSs on the conjugative transfer of ARGs, complementing current knowledge of the biotoxicity and potential ecological risks associated with BPNSs.202539827804
542130.9642Role of Yops and adhesins in resistance of Yersinia enterocolitica to phagocytosis. Yersinia enterocolitica is a pathogen endowed with two adhesins, Inv and YadA, and with the Ysc type III secretion system, which allows extracellular adherent bacteria to inject Yop effectors into the cytosol of animal target cells. We tested the influence of all of these virulence determinants on opsonic and nonopsonic phagocytosis by PU5-1.8 and J774 mouse macrophages, as well as by human polymorphonuclear leukocytes (PMNs). The adhesins contributed to phagocytosis in the absence of opsonins but not in the presence of opsonins. In agreement with previous results, YadA counteracted opsonization. In every instance, the Ysc-Yop system conferred a significant level of resistance to phagocytosis. Nonopsonized single-mutant bacteria lacking either YopE, -H, -T, or -O were phagocytosed significantly more by J774 cells and by PMNs. Opsonized bacteria were phagocytosed more than nonopsonized bacteria, and mutant bacteria lacking either YopH, -T, or -O were phagocytosed significantly more by J774 cells and by PMNs than were wild-type (WT) bacteria. Opsonized mutants lacking only YopE were phagocytosed significantly more than were WT bacteria by PMNs but not by J774 cells. Thus, YopH, -T, and -O were involved in all of the phagocytic processes studied here but YopE did not play a clear role in guarding against opsonic phagocytosis by J774. Mutants lacking YopP and YopM were, in every instance, as resistant as WT bacteria. Overexpression of YopE, -H, -T, or -O alone did not confer resistance to phagocytosis, although it affected the cytoskeleton. These results show that YopH, YopT, YopO, and, in some instances, YopE act synergistically to increase the resistance of Y. enterocolitica to phagocytosis by macrophages and PMNs.200212117925
7911140.9641Biochar induced inhibitory effects on intracellular and extracellular antibiotic resistance genes in anaerobic digestion of swine manure. Distribution of intracellular (iARGs) and extracellular ARGs (eARGs) in manure anaerobic digestion (AD) process coupled with two types of biochar (BC and BP) were investigated. And the effects of biochar on the conjugation transfer of ARGs were explored by deciphering the interaction of biochar with bacterial stress responses, physiological metabolism and antibiotic resistances. Results showed that AD process could effectively remove all the detected eARGs with efficiency of 47.4-98.2%. The modified biochar (BP) with larger specific surface area (SSA) was propitious to decrease the absolute copy number of extracellular resistance genes. AD process could effectively remove iARGs by inhibiting the growth of host bacteria. The results of structural equation models (SEM) indicated that biochar put indirect influences on the fate of ARGs (λ = -0.23, P > 0.05). Analysis on oxidative stress levels, antioxidant capacity, DNA damage-induced response (SOS) response and energy generation process demonstrated that biochar induced the oxidative stress response of microorganisms and enhanced the antioxidant capacity of bacteria. The elevated antioxidant capacity negatively affected SOS response, amplified cell membrane damage and further weakened the energy generation process, resulted in the inhibition of horizontal transfer of ARGs.202235609652
6752150.9641Isolation of radiation-resistant bacteria without exposure to irradiation. Resistance to desiccation was utilized in the selection of highly radiation-resistant asporogenous bacteria from non-irradiated sources. A bacterial suspension in phosphate buffer was dried in a thin film at 25 degrees C and 33% relative humidity. Storage under these conditions for 15 days or more reduced the number of radiation-sensitive bacteria. Further selection for radiation-resistant bacteria was obtained by irradiation of bacteria on velveteen in the replication process, thereby avoiding the toxic effect of irradiated media. The similarity of radiation resistance and identifying characteristics in irradiated and non-irradiated isolates should allay some concerns that highly radiation-resistant bacteria have been permanently altered by radiation selection.1979394680
575160.9640Identification and characterization of uvrA, a DNA repair gene of Deinococcus radiodurans. Deinococcus radiodurans is extraordinarily resistant to DNA damage, because of its unusually efficient DNA repair processes. The mtcA+ and mtcB+ genes of D. radiodurans, both implicated in excision repair, have been cloned and sequenced, showing that they are a single gene, highly homologous to the uvrA+ genes of other bacteria. The Escherichia coli uvrA+ gene was expressed in mtcA and mtcB strains, and it produced a high degree of complementation of the repair defect in these strains, suggesting that the UvrA protein of D. radiodurans is necessary but not sufficient to produce extreme DNA damage resistance. Upstream of the uvrA+ gene are two large open reading frames, both of which are directionally divergent from the uvrA+ gene. Evidence is presented that the proximal of these open reading frames may be irrB+.19968955293
8497170.9640Conjugation-mediated transfer of antibiotic resistance genes influenced by primary soil components and underlying mechanisms. Soil is the main natural reservoir of antibiotic resistant bacteria and antibiotic resistance genes (ARGs). Their dissemination and proliferation were largely motivated by conjugative transfer, while the influence of soil components on bacterial conjugative transfer and the underlying mechanisms remain poorly understood. In the present study, two Escherichia coli strains were exposed to soil minerals (quartz, kaolinite and montmorillonite) and organic matters (humic acid, biochar and soot) respectively to investigate their impact on ARGs conjugation. The results showed that quartz had no significant effect on conjugation; montmorillonite promoted the growth of the donor, but inhibited the recipient and conjugant; kaolinite and three organic matters significantly promoted the production of conjugant, while biochar promoted and then inhibited it with time prolong. Within the range of bacterial concentration involved in this study, the concentration of conjugant increased with the ratio of the concentration of donor and recipient (R(D/R)), indicating that the variation of conjugant production was mainly mediated by changing R(D/R). Further observation of biochar treatment group showed that the bacterial responses such as cell membrane permeability, cell surface hydrophobicity and biofilm formation ability shifted with the exposure time, which might be a potential factor affecting conjugative transfer. Collectively, our findings suggest that the type and exposure time of soil components jointly affected conjugation, while the change of R(D/R) and related bacterial responses are the main underlying mechanisms.202336586689
7985180.9639Differential response of nonadapted ammonia-oxidising archaea and bacteria to drying-rewetting stress. Climate change is expected to increase the frequency of severe drought events followed by heavy rainfall, which will influence growth and activity of soil microorganisms, through osmotic stress and changes in nutrient concentration. There is evidence of rapid recovery of processes and adaptation of communities in soils regularly experiencing drying/rewetting and lower resistance and resilience in nonadapted soils. A microcosm-based study of ammonia-oxidising archaea (AOA) and bacteria (AOB), employing a grassland soil that rarely experiences drought, was used to test this hypothesis and also whether AOB were more resistant and resilient, through greater tolerance of high ammonia concentrations produced during drought and rewetting. Treated soils were dried, incubated for 3 weeks, rewetted, incubated for a further 3 weeks and compared to untreated soils, maintained at a constant moisture content. Nitrate accumulation and AOA and AOB abundance (abundance of respective amoA genes) and community composition (DGGE analysis of AOA amoA and AOB 16S rRNA genes) were poorly adapted to drying-rewetting. AOA abundance and community composition were less resistant than AOB during drought and less resilient after rewetting, at times when ammonium concentration was higher. Data provide evidence for poor adaptation of microbial communities and processes to drying-rewetting in soils with no history of drought and indicate niche differentiation of AOA and AOB associated with high ammonia concentration.201425070168
8496190.9639Neglected resistance risks: Cooperative resistance of antibiotic resistant bacteria influenced by primary soil components. Various antibiotic resistant bacteria (ARB) can thrive in soil and resist such environmental pressures as antibiotics through cooperative resistance, thereby promoting ARB retention and antibiotic resistance genes transmission. However, there has been finite knowledge in regard to the mechanisms and potential ecological risks of cooperative resistance in soil microbiome. In this study, soil minerals and organic matters were designed to treat a mixture of two Escherichia coli strains with different antibiotic resistance (E. coli DH5α/pUC19 and E. coli XL2-Blue) to determine how soil components affected cooperative resistance, and Luria-Bertani plates containing two antibiotics were used to observe dual-drug resistant bacteria (DRB) developed via cooperative resistance. Results showed quartz, humic acid, and biochar promoted E. coli XL2-Blue with high fitness costs, whereas kaolin, montmorillonite, and soot inhibited both strains. Using fluorescence microscope and PCR, it was speculated DRB could resist the antibiotic pressure via E. coli XL2-Blue coating E. coli DH5α/pUC19. E. coli DH5α/pUC19 dominated cooperative resistance. Correlation analysis and scanning electron microscope images indicated soil components influenced cooperative resistance. Biochar promoted cooperative resistance by increasing intracellular reactive oxygen species, thereby reducing the dominant strain concentration required for DRB development. Kaolin inhibited cooperative resistance the most, followed by soot and montmorillonite.202235074748